Nonhost plant species as donors for resistance to pathogens with narrow host range I. Determination of nonhost status

Summary In crop species where disease and pest problems are serious, a common strategy is to attempt to introduce genes for resistance from more or less related (wild) species by means of interspecific crosses. Little attention has been given to the question, whether such a donor species should be a host or a nonhost if the target pathogen is a highly specialized micro-organism.In this paper it is discussed that, apart from obvious host and obvious nonhost relationships, various intermediate categories are to be discerned.Aspects like small numbers of accessions, taxonomic problems, environmental conditions and the age of the inoculated plants further hamper clear-cut verdicts whether a plant species is a host or nonhost.It is argued that, at least in the case of powdery mildew and rust fungi, histological observations are helpful in determining whether a predominant resistance of a plant species is based on avoidance, on pre-haustorial or on post-haustorial defence mechanisms. The former two mechanisms are typical of nonhost relationships, the latter one is the predominant mechanism of major genic host resistance, although exceptions occur. The questions for plant breeders would be which mechanism would provide the most effective protection of the crop and which would be better suitable for transfer to crop species.     

Analysis of disease resistance and quality characters of F1 hybrids of crosses between wheat (Triticum aestivum) and spelt (Triticum spelta)

Summary With the objective of creating new combinations of disease resistance and quality, hybrids between wheat and spelt (spelt is well adapted to cool and wet conditions) were produced and the expression of heterosis was analysed. Three winter wheat varieties were crossed reciprocally with two spelt cultivars and the F₁ hybrids were tested under artificial inoculation with stripe rust, powdery mildew and leaf rust. Disease susceptibility and quality characters (protein content, Zeleny value, grain hardness) were assessed in two year field trials. For stripe rust the F₁ hybrids were resistant if one of the wheat parents was resistant. Combinations with the susceptible wheat cultivar Arina were all susceptible irrespective of using a resistant spelt partner. Although the infection with powdery mildew was rather low, a similar reaction was obtained with the susceptible wheat variety Bernina. Leaf rust revealed very specific varietal influences. The two susceptible wheat varieties Bernina and Arina resulted in susceptible F₁ hybrids when combined with a moderately resistant spelt. Only when they were crossed with a resistant spelt cultivar the F₁ hybrids were resistant. Forno, a leaf rust resistant wheat, gave resistant F₁ hybrids in all combinations. Without exception the quality characters tested showed a negative heterosis effect resulting in protein levels and Zeleny values close to or below the values of the lower parent. It appears to be possible to produce resistant F₁ hybrids, mostly dominated by the resistance level of the wheat partner. The quality of the hybrids is mainly suitable for biscuit and spelt specific products; it needs specific screening for combinations with acceptable breadmaking quality.Abbreviations LSD Least Significant Difference - RH Relative Heterosis     

Accumulating polygenes for partial resistance in barley to barley leaf rust,Puccinia hordei. II. Field evaluation

Summary Eight lines from the cross between Vada and Cebada Capa with long to very long latent periods and four barley cultivars representing the known range of partial resistance to barley leaf rust, caused by Puccinia hordei, were evaluated in the field for partial resistance and in the greenhouse for the latent period (LP) in the young flag leaf.Each of the 12 entries was sown (15-4-1983) on a plot of 1.0 m². There were four replicates. To reduce interplot interference the plots were separated from each other by 4.0 m of spring rye. The number of urediosori per tiller was evaluated at 27-6, 4-7, 12-7 and ten days after heading. The LP was measured on 10 to 15 plants per entry in 1982 and on 10 plants in 1983.The levels of partial resistance varied greatly. The difference in number of sori per tiller between the most susceptible cultivar, Akka, and the most resistant cultivar, Vada, was about 50 times. Between Akka and the most resistant line this was approximately 5000 times. The LP's varied similarly. Vada had a LP 64% longer than that of Akka, the LP of line 26-6-11 was 15% longer. The range of partial resistance has been extended more than twofold.The correlation coefficients between LP and the level of barley leaf rust, expressed in transformed scale units, varied from -0.99 for the first sampling date to -0.97 for the third sampling date. Sampling the same development stage, ten days after heading, did not improve the r-value (r=–0.98). The LP evaluated in the young flag leaf is shown to be a very reliable criterion for partial resistance in the barley-Puccinia hordei pathosystem.Earliness tends to be associated with susceptibility. The correlation of days to heading with LP was 0.63, and with the level of barley leaf rust in the field 0.64.     

Genetics and histology of powdery mildew resistance in apple

Summary Eight scab-resistant cultivars and selections along with eight commercial apple cultivars were evaluated for powdery mildew resistance in greenhouse and nursery tests. Dayton, Liberty, Delicious and Tolman Sweet were rated moderately resistant to infection in both greenhouse and nursery tests. Segregation of seedlings among 14 progenies for mildew reaction indicated that mildew resistance is polygenically controlled in this material with additive gene effects. Recovery of mildew resistant seedlings from crosses involving a scab-resistant parent(s) suggested that this material can be useful in developing scab- and mildew-resistant apple cultivars. Histological investigations were conducted to describe mildew symptoms of infected leaves.     

Inheritance of resistance to Didymella lycopersici Klebb. in tomato (Lycopersicon Mill.)

Summary The inheritance of the resistance to Didymella lycopersici was studied on F₃- and Bc₁-lines from interspecific crosses of L. esculentum with L. hirsutum and with L. hirsutum glabratum. The resistance is not monogenic and is inherited in dominant fashion. The high h² values based on line means offer possibilities to seleet efficiently for enhanced levels of resistance.     

The concept of breeding for uniform and differential resistance and their integration

Summary It has been reported why resistance is needed in crop plants, how it has been searched for, and how long the resistance could be maintained in the host varieties. Uniform and differential resistance have been discussed and integrated within one genetic system. The possible evolution of both polygenes and major genes for resistance is discussed. Polygenes probably are older and produce the materials necessary for a host to show resistance, major genes acting as switching genes. Breeding for disease resistance has been briefly debated in connection with the discussion on host resistance.     

Determining levels of resistance to Tulip Breaking Virus (TBV) in tulip (Tulipa L.) cultivars

Summary The resistance of seven tulip cultivars to tulip breaking virus (TBV) was studied after mechanical inoculation by determining the number of infected plants and the level of virus in these plants over a period of one and a half years. Three cultivars (Apeldoorn, Kees Nelis and Lustige Witwe) appeared to be susceptible. Four cultivars (Cantata, Juan, Madame Lefeber and Princeps) showed a high degree of resistance. The number of infected plants of the latter cultivars was low and virus could not always be detected in the plants at the different sampling dates. Virus could not be detected in these plants after storage. These results indicate that the virus may disappear from or was not detectable in these plants. The levels of infection found are discussed with respect of its significance in studies of TBV in tulips.     

Components of partial resistance to Leptosphaeria nodorum among seven soft red winter wheats

Summary Seven soft red winter wheat cultivars were evaluated for partial resistance to Leptosphaeria nodorum under field conditions. The results demonstrate that resistance is available among cultivars that are adapted to the southeastern U.S. and that resistance is long lasting. Resistant cultivars had longer incubation and latent periods, slower rates of lesion development, and reduced the level of sporulation of L. nodorum. Seedlings of susceptible cultivars sustained severe disease with dew periods as short as 48 hr in greenhouse tests. One resistant cultivar sustained little damage with dew period as long as 144 hr.Latent period was shortest on the second leaf below the flag leaf (F-2 leaf) of all cultivars and longest on the flag leaf which reflected the effect of microclimate and leaf age. The range in length of latent period on the flag, F-1, and F-2 leaves of resistant cultivars was less than that for susceptible cultivars. Differences between cultivars were greatest (up to 6.8 days) for the F-2 leaf. A delay in production of inoculum on the F-2 and lower leaves of resistant cultivars should delay infection of the flag leaf and spike.The greatest differentiation among cultivars for sporulation of L. nodorum was on upper leaves at Feekes growth stage 11.2. Oasis consistently had less sporulation than other cultivars at all sampling dates and leaf positions. The rate of disease progress up the plant (disease severity) and area under the disease progress curve were also least on Oasis.There were significant correlations among components of resistance and associated components suggesting that a single or interrelated mechanisms control expression of resistance. Overall, Oasis was the most resistant cultivar followed by Coker 762 and Coker 747. There cultivars have remained resistant for ten or more years. Stacy was intermediate in resistance and was the most variable of the seven cultivars for the components tested. Holley, Omega 78, and Florida 301 were highly susceptible by all criteria tested. Plant height was not a factor in resistance. The resistant cultivars are mostly later in maturity than the susceptible cultivars. In this study, crop maturity was similar under existing environmental conditions during the two seasons when most data were collected.     

Stability of forage yield and quality in pearl millet hybrids heterogenous for rust resistance

Summary The susceptible pearl millet hybrid Tifleaf 1, the resistant hybrid Tifleaf 2, and two experimental 3-way hybrids with different proportions of resistant and susceptible plants were evaluated for rust resistance and forage yield and quality in 1990, 1991, and 1992. Different environmental conditions were obtained by varying planting date, planting density, and fungicide applications across three years of evaluation in the field. Rust severity of forage was negatively correlated with late season green yield, dry matter yield, in vitro dry matter digestibility, and digestible dry matter yield. The stability across environments of the 3-way hybrids was intermediate between resistant Tifleaf 2 and susceptible Tifleaf 1 for rust severity and the yield and quality measurements negatively correlated with rust severity. Mixtures of resistant and susceptible plants would probably provide greater control of rust than measured in these small-plot experiments. Although a mixture of resistant and susceptible plants appears to provide an alternative to monogenic control of rust in forage pearl millets, identification and utilization of additional sources of resistance would be beneficial to improve the performance of 3-way hybrids.     

The evaluation of adult plant resistance to powdery mildew (Erysiphe graminis f.sp. avenae) in transgressive lines of oats

Summary Previously identified segregant lines of oats with levels of adult plant resistance to powdery mildew (Erysiphe graminis f.sp. avenae) better than the resistant parent maintained this high level of resistance in field nurseries over two growing seasons. This enhancement was not expressed on inoculated detached leaves under laboratory conditions where no differences between the more resistant parent cv. Maldwyn and the most resistant segregants was detected. Reduced resistance was not detected in tests where leaf segments of the transgressive lines were inoculated with trained isolates. Problems associated with selecting for adult plant resistance under both field and laboratory conditions are discussed.     

Influence of race and post infection temperature on two components of partial resistance to wheat leaf rust in seedlings of wheat

Summary Components of partial resistance, infection frequency and latency period, were determined in 71 winter and spring wheat genotypes in the seedling stage, after infection with three races of leaf rust (Felix 3B, Clement B and Betuwe 85C) at three different day/night temperature regimes (24/18°C, 18/12°C and 12/6°C). The genotypes were split into two groups and two separate experiments were carried out. Five genotypes, SVP 84039, Akabozu, Banco, BH 1146 and Orso, conferred a low infection frequency and a long latency period and Westphal 12A a long latency period, indicating a relatively high level of partial resistance. The correlation coefficient between infection frequency and latency period was low. Race-specificity was not found. There was a significant temperature effect on the latency period. In the second experiment the temperature x genotype interaction was significant. Temperature-response functions of transformed data demonstrated that the latency periods of four relatively resistant genotypes, Westphal 12A, Banco, BH 1146 and Orso and of Sarno and Mirela were most sensitive to temperature. The range between the genotypes with the longest and the shortest latency period was highest at 12°C. Therefore, low temperature regimes are preferred to distinguish differences in level of partial resistance.     

Chromosome location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L.) 1. Mlk and other alleles at the Pm3 locus

Summary Several wheat cultivars/lines were inoculated with isolates of Erysiphe graminis tritici to identify new genes/alleles for resistance. The wheats were tested with 13 isolates that had been characterized from responses on differential lines with known resistance genes. Gene Mlk which occurs in cultivars Kolibri, Syros, Ralle and several other European common wheats was found to be an allele at the Pm3 locus and is now designated Pm3d. The mildew resistance in an old Australian wheat, W150, is conferred by a single gene also allelic to Pm3 and now designated Pm3e. The near-isogenic line Michigan Amber/8^*Cc possesses another allele now designated Pm3f. A Syrian land variety of common wheat shows mildew resistance that is conditioned by the combination of genes Pm1 and Pm3a. Finally, two accessions of Triticum aestivum ssp. sphaerococcum appeared to possess the Pm3c allele.     

Effect of the black sigatoka resistance locus bs 1 and ploidy level on fruit and bunch traits of plantain-banana hybrids.

Summary Plantain (Musa spp., AAB group) cultivation is threatened by black sigatoka, an airborne fungal (Mycosphaerella fijiensis Morelet) leaf spot disease. Several traits in plantains and bananas are mainly affected by major genes. The host resistance response to black sigatoka is under the control of at least three different genes, one major recessive gene bs ₁ and two other independent additive alleles. Diploid and tetraploid plantain hybrids were evaluated for bunch weight, fruit weight, fruit length and fruit circumference. The F₁ euploid hybrids were derived from interspecific crosses between the resistant diploid wild banana Calcutta 4 and the susceptible triploid plantain cultivars Obino l'Ewai and Bobby Tannap. Linear and multiple regression models, coefficients of determination, and Durbin-Watson statistics were used to determine the single and combined effects of the major locus for black sigatoka resistance and ploidy on the different traits in the progenies. Differences in yield were mainly due to changes in weight and girth of fruit, which are affected by black sigatoka disease. The combined effect of ploidy and resistance to black sigatoka was partially responsible for the quantitative trait variation in yield. As a result of the gene interaction in the black sigatoka resistance locus bs ₁, the partially resistant and less susceptible phenotypes showed higher yield than their more susceptible full sibs.     

An analysis of genes for resistance against Indian stem rust races in two bread wheat cultivars

Summary The genetic constitution of two bread wheat accessions from the International Spring Wheat Rust Nurseries (E 5883 and E 6032) has been studied for reaction to four Indian races of stem rust. Analysis of E 5883 has revealed that for each of the races 15C, 21 and 40 a single dominant gene operates for resistance. The dominant gene against race 15C was identified as Sr6. The dominant genes for resistance against races 21 and 40 were found to be different from the genes described so far. Resistance against race 122 is controlled by a single recessive gene producing characteristically a 2 type of reaction. This gene was identified as Sr8.The resistance of E 6032 against each of the races 15C, 21 and 40 is controlled by two genes, one dominant and one recessive, which act independently. Dominant genes effective against 15C, 21 and 40 were conclusively identified as Sr6, Sr5 and Sr9b, respectively. From the correlated behaviour against races 15C and 40 as well as from the phenotypes of the resistance reactions rhe same recessive gene, undescribed so far, operates against the two races. The second recessive gene operating against race 21 was also observed to be different from those so far designated. E 6032 was, however, found to be susceptible to races 122.The presence of Sr6 both in E 5883 and E 6032 against race 15C was further confirmed through F₂ and F₃ segregation data.     

Genetic analysis of resistance to Phoma tracheiphila in three Citrus and Poncirus progenies

Mal secco is a serious tracheomycosis, caused by the fungus Phoma tracheiphila (Petri) Kanc, et Ghik., which attacks some species of the genus Citrus. The considerable amount of time and space required to evaluate host plant response has discouraged research on the mechanism of inheritance involved in the susceptibility to this disease. The aim of the present work was to study segregation of the resistance trait in three progenies obtained by crossing Citrus latipes (Swing.) Tan., as a female parent, with the male parents Poncirus trifoliata (L.) Raf., C.aurantium L. and C. volkameriana Pasq. Resistance determination was based on natural infections observed over 10 years. Results allowed us to hypothesize three alternative genes (A, B and C) which determine dominant resistance as opposed to susceptibility: the presence of a single allele can thus confer resistance. In addition, we suggest the existence of a fourth gene which, in conditions of dominance, is able to determine susceptibility by nullifying the action of the dominant B allele. Response to the pathogen was also associated with the production of chitinase a PR (pathogenesis-related) protein reported to be present in great quantities in mal secco-resistant Citrus genotypes. Segregation analysis of one F₂ progeny confirmed the hypothesis that the resistance trait is controlled by at least two dominant genes. Further evidence is needed to ascertain whether this hypothesis is applicable to other Citrus species, in which the possibility of other genetic mechanisms being implicated cannot be ruled out, thus making the genetic basis of mal secco resisance more complex.     

Genetic analysis of inbred lines and their crosses for resistance to head blight (Fusarium culmorum,F. graminearum) in winter rye

Summary For genetic analysis of head blight in winter rye (Secale cereale) caused by Fusarium culmorum, six homozygous inbred lines from the Petkus gene pool were crossed in all combinations to obtain 15 diallel F₁ crosses and the corresponding 15 F₂ crosses. These materials and 10 additional inbreds were artificially inoculated in a 2-year field experiment. The inbreds were also tested with F. graminearum in a separate sub-experiment.Single disease rating, average disease rating, and yield components (grain-weight per spike, 1000-grain weight, kernel number per spike) relative to the non-inoculated treatment were significantly affected by Fusarium head blight in all material groups. The relative grain weight per spike ranged from 26% to 88%. Significant genotypic and genotype x year interaction variances were found throughout. Heritabilities were highest for homogeneous inbreds (h²=0.6–0.8) and lowest for heterogeneous F₂ crosses (h²=0.4–0.6). Disease rating and relative grain-weight per spike were highly correlated for the inbreds and F₂ crosses (r0.7, P0.01), but lower for the F₁ crosses (r0.6, P0.05). Inter-annual correlation coefficients for disease ratings and relative grain-weight per spike ranged from r0.7 (inbreds) to r0.5 (F₂ crosses). The diallel analysis showed significant GCA effects only for relative 1000-grain weight in 1990, but significant SCA and SCAx year interaction variances for most traits. The resistances of 16 inbreds to F. culmorum and F. graminearum were tightly associated for all traits (r=0.96–0.97, P0.01).In conclusion, only slow progress can be expected from selecting for Fusarium head blight resistance in rye due to the limited amount of additive genetic variance and the great improtance of environmental factors.     

Histological studies on the infection of triticale,wheat and rye byPuccinia recondita f.sp.tritici andP. recondita f.sp.recondita

Summary The reaction of eight triticales and of the respective wheat and rye parental lines to infection by the leaf rust fungi of wheat and rye were studied in the seedling stage. The histological observations indicated that wheat and triticale showed a typical nonhost reaction to the leaf rust of rye: sporelings of this fungus were arrested after the formation of primary infection hyphae and before the formation of extensively branched mycelium, mostly without necrosis of plant cells. The rye inbred lines were all susceptible to the rye leaf rust. The reaction of wheat and triticales to the wheat leaf rust was susceptible or resistant. The reaction of resistant lines could be early or late and complete or incomplete, but was associated with substantial necrosis of plant cells, and therefore entirely different from the nonhost reaction to rye leaf rust. In their reaction to wheat leaf rust the rye lines were similar to the resistant wheat and triticale lines. They did not show an important degree of nonhypersensitive early abortion as would be expected in a nonhost species. It appeared that genes for hypersensitive resistance in triticale may be contributed by either the wheat or the rye parental line.A screening of sixty wheat, rye and triticale lines confirmed the nonhost status of wheat and triticale to rye leaf rust and the hypersensitive or moderately susceptible reaction of rye to wheat leaf rust.     

Breeding for resistance to yellow disease of hyacinths. II. Influence of flowering time,leaf characters,stomata and chromosome number on the degree of resistance

Summary Analysis of an incomplete diallel design with 14 hyacinth cultivars and 4 wild accessions of Hyacinthus orientalis showed a significant GCA component for resistance to yellow disease. The late flowering hyacinth cultivars King of the Blues and Marconi and the early flowering Hyacinthus orientalis 70129 were found to be the best combiners for yellow disease resistance. Within a cultivar, the degree of earliness itself was not associated with the degree of resistance. Correlation coefficients of leaf characters and degree of resistance showed that hyacinths with short and/or narrow leaves are generally more resistant than those with long and/or broad leaves. Tetraploid cultivars with few, large stomata tend to be susceptible, diploid ones with many small stomata tend to be the more resistant.     

Allelism of genes controlling resistance to TMV in Capsicum L.

Summary The genes for resistance to pepper strains of TMV in five C. chinense Jacq. accessions, the PI numbers 152225, 159236, 315008, 315023 and 315024, were investigated for allelism. The resistance of these five C. chinense accessions was found to be inherited monogenically and to be partially dominant. The resistance genes in these accessions appeared to be allelic and were also found to be allelic with the known alleles L¹ and L. It is proposed to use the symbol L³ for these resistance genes.     

Mapping of leaf and neck blast resistance genes with resistance gene analog, RAPD and RFLP in rice

An F₈ recombinant inbred population was constructed using a commercial indica rice variety Zhong 156 as the female parent and a semidwarf indica variety Gumei 2 with durable resistance to rice blast as the male parent. Zhong 156 is resistant to the fungus race ZC₁₅ at the seedling stage but susceptible to the same race at the flowering stage. Gumei 2 is resistant to ZC₁₅ at both stages. The blast resistance of 148 recombinant inbred lines was evaluated using the blast race ZC₁₅. Genetic analysis indicated that the resistance to leaf blast was controlled by three genes and the presence of resistant alleles at any loci would result in resistance. One of the three genes did not have effects at the flowering stage. Two genes, tentatively assigned as Pi24(t) and Pi25(t), were mapped onto chromosome 12 and 6,respectively, based on RGA (resistance gene analog), RFLP and RAPD markers. Pi24(t) conferred resistance to leaf blast only, and its resistance allele was from Zhong 156. Pi25(t) conferred resistance to both leaf and neck blast, and its resistance allele was from Gumei 2. In a natural infection test in a blast hot-spot, Pi25(t) exhibited high resistance to neck blast, while Pi24(t) showed little effect. This revised version was published online in August 2006 with corrections to the Cover Date.